People often think of ecosystems in terms of the plants and animals that live in them, but a lot of other factors, like geology, altitude, and climate, are crucial to determining what kind of ecosystem develops in a particular place. When one of those non-living factors changes, ecosystems change, too.

Over the 20th century, global temperatures increased by an average of about 1.3o F, but some places have warmed a lot more than this, and other places have warmed less. These temperature increases have been enough to trigger changes in ecosystems all over the world, especially in places where the warming has been the greatest. In some places the changes have been subtle, like a slight shift in vegetation that only a careful observer would notice. In other cases small changes in climate have sparked a chain of larger effects, leading to massive changes.

The biggest climate-caused ecosystem shifts today are happening at the world's most northern latitudes, where the temperature over the last century has been rising about two times faster than the global average. In Alaska, for example, warming has paved the way for a spike in the numbers of spruce bark beetles. Bark beetles have been a pest to Alaskan white spruce for thousands of years, but their numbers were held in check by the cold climate, which forced the insects to hole up in the bark of a individual trees for most of the year. As the length of the warm season increased over the 1980s and 1990s, however, bark beetles had more time to fly from one tree to the next, burrow, and lay their eggs between the bark and wood. The beetles had another thing going for them, too: A multi-year drought had weakened many of the spruce trees, leaving them vulnerable to attack. In the mid-1990s the bark beetle population exploded, and over the next few years the pests wiped out white spruce forests over an area the size of Connecticut. In the years since, the combined forces of a longer insect breeding season and forest management practices that left forests overcrowded gave way to similar epidemics further south. Large swaths of pine and spruce have been destroyed by insects in New Mexico, Colorado, Wyoming, and elsewhere.

In the late 1990s the effects of the bark beetle epidemic rippled throughout Alaska's white spruce ecosystem and affected virtually every population of living organisms -- but not all of the impacts were negative. Fewer spruce trees meant a sunnier understory in the forest, which allowed grasses to move in and take hold. The grasses, in turn, changed the soil temperature, making the environment more friendly for some other types of vegetation. Animals who feed on grasses, including moose, elk, and some birds, also benefited.

But the beetle infestation was bad news for organisms that rely on white spruce for their habitat, like hawks, owls, red squirrels, and voles. Voles, a type of small, round rodent, are an especially vital part of the ecosystem because they help spread mycorrhizal fungi, which attaches to the roots of plants and helps them take in water and nutrients. Voles are also an important food for a number of predators.

Because ecosystem changes always hurt some living creatures and help others, it's hard to say whether a change is good or bad overall. Instead, people who study ecosystems often focus on the impacts on a single species -- for instance, us. In the short term the Alaskan spruce beetle epidemic supplied a lot of people with firewood, but only by destroying tons of otherwise valuable timber and threatening the livelihoods of loggers. And no one knows for sure what the long-term impacts on the forest will be. Ecosystems tend to return to their previous states after disturbances like pest outbreaks, fires, or major storm events, but if the Alaskan spruce ecosystem is disturbed too often or too much, it might shift to a different type of forest, a woodland, or a grassland instead.

In extreme cases major assaults on ecosystems can lead to a total collapse, in which the ecosystem doesn't bounce back to the way it was or transition to a new, healthy state. The result is an area with very little life; in the oceans, biologists refer to these areas as "dead zones." One example of ecosystem collapse is the coral reef die-off in the Indian Ocean in the late 1990s. The reefs had been up against multiple pressures for several decades, including reduced populations of fish (a result of overfishing), ocean acidification due to increased CO2, and rising water temperatures. Then, in 1998, a strong El Niño event moved warmer-than-usual water into the coral reefs, causing massive coral bleaching. As a result, one quarter of the coral in the Indian Ocean died, taking with it much of the marine life that depended on it for survival. Eight years later researchers reported that in some places in the Indian Ocean, the number of fish species has been reduced by half. Other coral reef collapses occurred in parts of the Caribbean in 2005 and 2010, also due to extreme heat events.